Spatial heterogeneity and biotic interactions – scaling from experiments to natural systems

Much of current ecological theory stems from experimental studies. These studies have often been conducted in closed systems, at spatial scales that are much smaller than the systems of interest. It is known that the outcome of these experiments may be seriously affected by artefacts associated with the caging procedures, as well as by the actual difference in spatial scale between experimental and target system. Yet, quantitative methods for estimating and removing artefacts of enclosure and for extrapolating experimental results to the scales of natural systems are largely lacking. The aim of this thesis was to confront some of the problems encountered when scaling from experiments to nature in studies on predator-prey systems, with focus on effects of changes in spatial heterogeneity. Specifically, I examined mechanisms that may cause consumption rate estimates to depend on the size of the experimental arena. I also studied methods for scaling up these process rate estimates to natural predator-prey systems. The studies were performed on invertebrate predator-prey systems found in the northern Baltic Sea. Initially, a descriptive study of small-scale distribution patterns was performed, in order to get background information on how the behaviour of the organisms was manifested in the spatial structure of the community. Experimental studies of two predator-prey systems exposed an artefact that may be widespread in experiments aiming at quantifying biotic interactions. It is caused by predator and prey aggregating along the walls of the experimental containers. This behaviour affects the encounter rate between predator and prey, thereby causing consumption rates to be scale-dependent. Opposing the common belief that larger arenas always produce less biased results, this scale effect may instead be reduced by decreasing arena size. An alternative method for estimating the magnitude of, and subsequently removing, the artefact caused by aggregation along the arena wall was presented. Once unbiased estimates of process functions have been derived, the next step is to scale up the functions to natural systems. This extrapolation entails a considerable increase in spatial heterogeneity, which may have important implications for the dynamics of the system. Moment approximation provides a method of taking the heterogeneity of natural populations into account in the extrapolation process. In the last study of the thesis, the concepts of moment approximation and how to estimate relevant heterogeneity were explained, and it was shown how the method may be used for adding space as a component to a dynamic predator-prey model. It was concluded that moment approximation provides a simple and useful technique for dealing with effects of spatial variation, and that a major benefit of the method is that it provides a way of visualising how heterogeneity affects ecological processes.